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The Affinities of the Rat-Kangaroos (Marsupialia) As Revealed by a Comparative Study of the Female Urogenital System

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The Affinities of the Rat-Kangaroos (Marsupialia) As Revealed by a Comparative Study of the Female Urogenital System PAP. & PROC. ROY. Soc. TASMANIA, 1945 (1ST OCTOBER, 194G) The Affinities of the Rat-kangaroos (Marsupialia) as revealed by a comparative study of the Female Urogenital System. BY JOSEPH PEARSON (Read 13th November, 1945) 1. INTRODUCTION According to the accepted view the family Macropodidae is divided into three sub-families, the Hypsiprymnodontinae, the Potoroinae, and the Macropodinae. The members of the first two are collectively known as the rat-kangaroos and the third sub-family comprises the true kangaroos and wallabies. The present paper is concerned mainly with the rat-kangaroos, which consist of five genera, as follows:- Sub-family HYPSIPRYMNODONTIN AE (1) Hypsiprymnodon Ramsay, 1876 Con.tains a single little-known species recorded from Queensland. This genus is the only form in which the hallux is present. Mainly for this reason it is regarded as the most primitive rat-kangaroo and is usually placed in a separate sub-family from the rest. Sub-family POTOROINAE (2) Potoroiis Desmarest, 1805 Three species have been recorded of which probably only one, P. t1·idactylus (Kerr, 1792), can now be obtained. ( 3) Bettongia Gray, 1837 There are four species, all of which are rare or extinct, with the exception of the Tasmanian form, B. cuniculus ( Ogilby, 1838). (4) Aepyprymnus Garrod, 1875 Contains only one species, A e. rufescens (Gray, 1837), which IS extremely rare. Nothing is known of its internal anatomy. ( 5) Calowymnus 0. Thomas, 1888 A single species, C. ca·mpestTis (Gould, 1843), which was re-discovered fifteen years ago after a lapse of ninety years. The female urogenital system of only two of these genera is known, viz., Bettongia (Owen, 1834; Brass, 1880; Pearson, 1944, 1945) and PotoToiis (Pearson, 1944, 1945). In the present paper descriptions of the female urogenital system of Caloprymnus and Hypsipryrnnodon are given. Nothing is known of the internal anatomy of Aepyp1·ymnus. 13 14 THE AFFINITIES OF THE RAT-KANGAROOS 2. THE FEMALE UROGENITAL SYSTEM OF CALOPRYMNUS CAMPESTRIS (Figs 1-3) Cctloprymnns cawpestris was first described by Gould in 1843 as Bettongict campestris from three specimens obtained by Sir George Gray. After a lapse of nearly ninety years a single specimen was obtained by Mr. Reese from the north­ eastern part of South Australia, and this important re-discovery led to a special expedition being sent out in 1931, with the result that Mr. Finlayson, Honorary Curator of Mammals, South Australian Museum, secured several specimens of this ra1·e species and made important observations on its habits, external structure, and probable affinities (Finlayson, 1932). Through the good offices of the Director of the National Museum, Melbourne, an opportunity has now been presented of reporting upon its female urogenital system, thereby extending our knowledge of the com­ parative anatomy of this important group of marsupials. This help so readily afforded is gratefully acknowledged. The following description of the female urogenital system of Caloprymnus is based upon an examination of spi1·it specimen No. R. 13609 from the collections of the National Museum, MelboUJ·ne. The parts described have been dissected but have not been detached from the carcass, and should be available, therefore, for future examination in Melbourne. The external measm·emeuts of the spirit specimen are as follows:- Length of head and body 34 em. Length of tail 33cm. Length of ear 3·7 em. Length of pes 11·2 em. The specimen examined is a mature female. The left uterus is considerably enlarged and contains a foetus which, though recognizable, is not sufficiently well preserved to enable the details of its structure to be made out. However, the urogenital system in general is in a satisfactory state of preservation and no difficulty has been experienced in making out the details which are given below. A general inspection of the female urogenital system shows that it resembles that of Potoroiis to a marked degree, though the anterior vaginal expansion is relatively much larger and bears out the suggestion made in previous papers (Pearson, 1944, 1!J45) that this portion of the vagina is, in fact, an incipient caecum such as is seen in full development in Bet.tongia. Uteri (l.L~t., r.ut.) The two uteri would, in normal circumstances, have a somewhat similar appear­ ance to those of Potoroiis as described by Pearson (1945), but, as mentioned above, the left uterus of this specimen is considerably enlarged owing to the presence of a foetus, and has a length of 47 mm. (body and neck) and a maximum width of 22 mm. The normal 1·ight uterus is 28 mm. long and 12 mm. wide. lVledian vaginal cnl-de-sae (m.e.) This has an extreme length of about 26 mm. and the first (anterior) quarter forms part of the anterim vaginal expansion. The cul-de-sac is about 5 mm. wide in its second qua1·ter and tapers gradually as it passes caudally. It ends blindly immediately ante1·ior to the junction of the two lateral vaginae. A ntcrior vagi1wl ccqJrtnsion ( ct.v.e.) This is a commodious, winged chamber for·med by the hypertrophy of the anterior portion of the vaginal system. It occupies much the same position as the comparable structure in Poturoiis, but differs from it in being larger and possessing two well­ defined spheroidal wings which are snmewhat flattened dorso-ventrally. The outlines JOSEPH PEARSON 15 of these wings are somewhat irregular, which-may be due, in part, to the effects of the preservative. This anterior vaginal expansion has a single continuous cavity, the medial portion of which may be regarded as the anterior part of the vaginal cul-de-sac already described. The lumen of each wing opens freely into this central cavity and also communicates posteriorly with the lumen of the contiguous lateral vagina. bl. !.~.~: w·eth. 1 pv.s. 3 red.-ct.-- 1~· dp.--- (Figs 1-3) Caloprym.nus campestris DiagTams of the female urogenital system. FIG. I.-Horizontal section, Dorsal view x lli. FIG. 2.-Horizontal section, Ventral view x lf,. FIG. 3.-Paramedial section x 1!. NoTE.-In all figures 1Jink represents the vaginal system; blue represents the urinary system. a.v.e.-A nterior vaginal expansion m.c.-Median vaginal cul-de-sac' (becoming a true bl.-Urinary bladder median vagina i:rl fig. 7) ct.-Clitoris p.v.s.-Posterior vagina] sinus d.p.-Digital process projecting from the postero- rect.-Rectum dorsal lip of the cloaca. r. ut.-Right uterus (normal size) l.l.v.-Left lateral vagina u.g.s.--Urogenital sinus I. ur.-Left ureter u.o.-Opening of urethra into urogenital sinus 1.ut.-Left uterus, considerably swollen and con­ ur.-Ureter taining a foetus. ureth.-Urethra l.v.-Lateral vagina. ut.-Uterus v.c.-Vagina] caecum. 16 THE AFFINITIES OF THE RAT-KANGAROOS The extreme width of the anterior vaginal expansion is about 27 mm. Each wing has an antero-posterior length of about 15 mm. and a width of about 11 mm. As in the case of Potoroiis the uterine necks project from the roof of the median portion of the chamber on papillae which are not so well developed as in Potoro'iis. As in that genus each papilla is perforated near its tip by the os uteri. Although there can be little doubt that the vaginal system is completely separated into right and left elements in early development, the septum separating the two halves completely disappears in the mature female. LateTal vaginae (l.v.) and posterioY vaginal sinus (p.v.s.) Each lateral vagina arises from the postero-lateral wall of the corresponding wing of the anterior vaginal expansion and has a length of about 21 mm. The right and left lateral vaginae have a straight course as they pass caudally and gradually converge to meet behind the blind extremity of the median cul-de-sac. They are widest near their anterior end, where they have a width of about 5 mm. and gradually narrow to a width of 3 mm. or less. The two coalesce to form the posterior vaginal sinus, which, as in the case of both Potoroiis and Bettongia, is a median tube lying immediately dorsal to the posterior section of the urethra. Both the posterior vaginal sinus and urethra open togethei· into the urogenital sinus. The posterior vaginal sinus is 8 mm. long and about 5 mm. wide. It is shorter than the comparable structure in Potor-oiis, but slightly longer than that of Bettongia: Urinary bladder (bl.) and urethra (ur-eth.) As in Poto·roiis and Bettongia the urinary bladder has an extreme anterior attachment situated immediately behind the anterior vaginal expansion (or caecum). The urethra runs ventral to the median cul-de-sac and posterior vaginal sinus, and together with the latter, opens into the urogenital sinus. The urethra has a total length of about 26 mm. It is about 3 mm. wide anteriorly and narrows somewhat towards its posterior end. Urogenital sinus (u.g.s.) This is about 14 mm. long and about 5 mm. wide. There is a well-developed clitoris attached to the ventral wall near the posterior extremity. 3. FEMALE UROGENITAL SYSTEM OF HYPSIPRYMNODON MOSCHATUS (Figs 4, 5.) After the present paper' had been completed in its original form an unpublished account by Miss F. R. Heighway of the anatomy of this species was brought to the writer's notice through the good offices of Professor Abbie of the Anatomy Depart­ ment, University of Adelaide. Later Professor Burkitt of the Anatomy Department, University of Sydney, in whose laboratory Miss Heighway's work was carried out, was kind enough to allow the writer to examine a dissection of the female urogenital system of this species which had been made by Miss Heighway in the course of her work.
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